EAGLE H, HABEL K
J Exp Med. 1956 Aug 1;104(2):271-87. doi: 10.1084/jem.104.2.271.
Only minimal amounts of poliomyelitis virus were formed by HeLa cells placed in a medium free from glucose and glutamine, even if the medium contained an otherwise full complement of essential and non-essential amino acids, purines, pyrimidines, NB(4) (+), and serum protein. Conversely, within one log of the optimal yield of virus was formed by HeLa cells in a medium containing only glucose, glutamine, and salts, even if the cells had been starved in this medium for 12 hours prior to their inoculation. The presence of glucose alone caused an average 170-fold increase in viral output beyond the amounts formed by the glucose- and glutamine-depleted cells. The addition of glutamine alone caused an average 2000-fold increase; and the addition of both increased the viral formation 40,000-fold. Qualitatively similar results were obtained with unstarved cells, not previously depleted of glucose and glutamine. It follows that only a small proportion of HeLa cells are capable of forming virus unless either glucose or glutamine, or both, are present in the medium. The elaboration of virus was significantly delayed in media containing glucose but no glutamine. The absence of glucose and glutamine did not prevent the fixation of poliomyelitis virus by the cell. When these compounds were added to previously depleted cells even 6 hours after inoculation, and after the excess free virus had been removed by washing and by the addition of specific antiserum, normal amounts of virus were formed despite the degenerative changes caused by the previous glucose and glutamine deprivation. Possible functions of glucose and glutamine in the elaboration of virus are discussed in the text. Such factors other than glucose, glutamine, or salts (e.g. amino acids, purines, pyrimidines, vitamins, protein, or NH(4) (+)) as may be needed by HeLa cells for the propagation of poliomyelitis virus, need not be present in the medium and cannot be easily washed out of the cell. Even 12 hours' total deprivation of the cells in salt solution prior to inoculation only slightly decreased their virus-synthesizing capacity in a similarly deficient medium, provided only that adequate amounts of glucose and glutamine were retained.
将HeLa细胞置于不含葡萄糖和谷氨酰胺的培养基中时，即使该培养基含有其他全部必需和非必需氨基酸、嘌呤、嘧啶、NB(4)(+)以及血清蛋白，也只会产生极少量的脊髓灰质炎病毒。相反，HeLa细胞在仅含葡萄糖、谷氨酰胺和盐的培养基中能产生接近最佳产量的病毒，即便这些细胞在接种前已在该培养基中饥饿培养12小时。仅葡萄糖的存在就能使病毒产量比缺乏葡萄糖和谷氨酰胺的细胞所产生的量平均增加170倍。仅添加谷氨酰胺能使病毒产量平均增加2000倍；同时添加二者则使病毒形成量增加40000倍。对于未饥饿处理、之前未耗尽葡萄糖和谷氨酰胺的细胞，也得到了定性相似的结果。由此可见，除非培养基中存在葡萄糖或谷氨酰胺，或二者都存在，否则只有一小部分HeLa细胞能够产生病毒。在含有葡萄糖但不含谷氨酰胺的培养基中，病毒的产生会显著延迟。缺乏葡萄糖和谷氨酰胺并不妨碍细胞对脊髓灰质炎病毒的吸附。即使在接种6小时后，甚至在通过洗涤和添加特异性抗血清去除过量游离病毒之后，将这些化合物添加到先前耗尽的细胞中，尽管先前的葡萄糖和谷氨酰胺缺乏导致细胞发生了退行性变化，但仍能形成正常量的病毒。文中讨论了葡萄糖和谷氨酰胺在病毒产生过程中的可能作用。HeLa细胞繁殖脊髓灰质炎病毒可能所需的除葡萄糖、谷氨酰胺或盐之外的其他因素（如氨基酸、嘌呤、嘧啶、维生素、蛋白质或NH(4)(+)），不一定存在于培养基中，也不容易从细胞中洗脱。即使在接种前将细胞在盐溶液中完全饥饿培养12小时，只要保留足够量的葡萄糖和谷氨酰胺，在同样缺乏营养的培养基中，其病毒合成能力也只会略有下降。
